E DEVELOPMENT OF THE PERITHECIA. IN THE 
MICROTHYRIACKAE AND A COMPARISON 
WITH MELIOLA 


BY 


RUTH WINIFRED RYAN 


AB. oaintaClara College,.1921 
A.M. University of Illinois, 1923 


THESIS 


Spiele euNee AR PLA FULFILLMENT OF THE REQUIREMENTS 
Pore een POP DOCTOR OF PHILOSOPHY IN BOTANY: 
iNet a OU ATE SCHOOLLOP. THE UNIVERSITY 
OF TIELINOIS, 2925 


Reprinted from Mycologia, Vol. XVIII, No. 3, May-June, 1926 


THE DEVELOPMENT OF THE PERITHECIA IN 
THE MICROTHYRIACEAE AND A COM- 
PARISON WITH MELIOLA 


[Reprinted from Mycotocia, Vol. XVIII., No. 3, May-June, 1926.] 


THE DEVELOPMENT OF THE PERITHECIA IN 
THE MICROTHYRIACEAE AND A COM- 
PARISON WITH MELIOLA 


RutTH WINIFRED RYAN 


(WitH PLates 12-15) 


In recent years many advances have been made in the tax- 
onomy of the fungi. Perhaps in no one group has there occurred 
as many changes as in the Microthyriaceae. Under the leader- 
ship of von Hohnel the entire bases of the family became subject 
to critical investigation followed by’ a complete revolution of 
the classification. 

The family was placed among the Dothideales by Saccardo (9) 
in 1883. Since then it has been described as belonging to the 
Perisporiales. Atkinson (2), however, considered that they have 
little in common with this order. He believed that the Micro- 
thyriaceae represent reduced forms derived on the one hand from 
the Phacidiales and, perhaps, on the other from the Sphaeriales. 
He did not say definitely, though, where they do belong. Theis- 
sen and Sydow (15) placed the family in a new order, the Hemi- 
spheriales, which they created for genera having a superficial, 
halbert- to shield-shaped perithecium. Miss Doidge (4) accepts 
this classification in her systematic work. According to Arnaud 
(1) the family belongs to the Pyrenomycetes, because, he says, 
the asci are localized in particular zones formed by the gelatiniza- 
tion of the sterile cells of the cavity. The Microthyriaceae 
would thereby be near neighbors of the Myriangiaceae. In my 
taxonomic study of the Microthyriaceae of Porto Rico (8) and 
Hawaii (10), I used the classification given by Theissen and 
Sydow. The Microthyriaceae seem to me more closely related 
to the Hemisphaeriales than to the other groups with which the 
family had formerly been placed. 

The first description for the family is given by Saccardo (9). 
Translated, it reads “‘simple perithecia superficial, black, mem- 
branous to carbonaceous, dimidiate, flattened; context nearly 


100 


RYAN: PERITHECIA IN MICROTHYRIACEAE 101 


always radiate.’’ This description is accurate. The characters 
are firmly established and easily recognizable. 

The various genera of the family can be placed in two groups 
according to Theissen (12) and Arnaud (1), those having a free 
mycelium and those in which it is lacking. The present paper 
deals with the development of the perithecia in the eleven genera 
that have a superficial mycelium. The material for study was 
obtained from the herbarium of the University of Illinois from 
collections made by Dr. F. L. Stevens in Porto Rico, Hawaii, 
and British Guiana. 

The mycelium in all the eleven genera plays an essential role 
in the development of the perithecium. Its characteristics are 
well determined and hereditarily fixed. It is brown, cylindrical, 
septate, tough and may or may not bear hyphopodia or nodulate 
cells. These last two structures also play a part in the formation 
of the perithecium. 

Formerly it was thought that the hyphopodia when they 
occurred on the mycelium of fungi might be sexual organs. This 
was at first supposed to be the case in Meliola. Ward (17) in 
studying this genus suggested that the hyphopodia were such. 
More recently Thaxter (11) has compared them to the antheridia 
of the Laboulbeniales. Arnaud (1) is of the opinion that they 
do not function as sexual organs and that they are merely the 
rudiments of the perithecia and absorptive organs. Investiga- 
tions by Maire (6) show that the hyphopodia in Meliola function 
as organs of absorption and attachment. Cytological studies by 
Arnaud (1) seem to indicate that the same relation to the host 
tissue exists in the Microthyriaceae. 

Arnaud includes Meliola in his study of the Asterinées (1). 
To me they seem fundamentally different as portrayed in their 
development. For this reason I have placed them in the Peri- 
sporiaceae as does Beeli (3). 

The hyphopodia in Melola are two-celled, while they are 
either one- or two-celled in the Microthyriaceae. In the Micro- 
thyriaceae and in Meliola they may be opposite or alternate, the 
majority being the latter. Often one finds several hyphopodia 
arranged on one side of the mycelium, while further on in the 
same filament they will be either opposite or alternate. The 


102 ; MYCOLOGIA 


characteristic arrangement of the structure may be used as an 
aid in classification. 

The formation of the perithecium in the majority of ascomy- 
cetes results from the activity of sexual organs, which have the 
power of conjugation. This fusion then is the point of develop- 
ment of the perithecium. In Melola (17) the hyphopodia sub- 
stitute for these organs. Those that give rise to perithecia have 
a reflexed swollen end and produce the fruiting structures through 
proliferation. In the Microthyriaceae the hyphopodia though 
not sexual organs give rise to the perithecia in certain cases. This 
family advances further, for here the perithecia may arise inde- 
pendently of the hyphopodia, that is, from the mycelial cells 
themselves. This method of origin seems to be by far the most 
common. 

A search through mycological literature revealed only a few 
articles which bear in any way on the development of these fungi. 
Gaillard (5) studied a few species of Asterina. He says that the 
summit of the hyphopodium turns toward the leaf. This small 
cell divides, at the same time becoming clearer. The primitive 
membrane becomes thinner, budding in places and appearing 
as a small lenticular body closely applied against the leaf. Its 
surface is covered by deep lobes, which in growing seem to radiate 
around a point, which is the place of insertion on the mycelium 
of the primitive hyphopodium. In the majority of cases the 
hyphopodia do not participate alone in the division, but the 
portion of the mycelium where it is inserted also takes an active 
part. Thus he indicates one method of development charac- 
teristic of the genus and of the family as a whole. 

Raciborski (7) refers to the subject only in passing, saying 
that the perithecia are borne laterally on the hyphae as in Balla- 
dyna, Aldona, Dimerosporium, or Melola, and that they grow 
conical and broaden against the leaf. Miss Doidge (4) briefly 
gives two methods; from a medial cell of a hypha and from the 
terminal cell of a short lateral branch. She does not go into 
any detail. 

Theissen (13) says that divisions in the hyphae give rise to 
irregular cells by the insertion of several cross walls. The 
perithecium arises on the lower surface of the hyphae. A small 


d 


RYAN: PERITHECIA IN MICROTHYRIACEAE 103 


knot of cells is formed and is the base of the perithecium, which 
continues to grow radially and centrifugally to a disk-shaped 
structure. The radial walls remain locked together. New radial 
walls become inserted so that the disk shape is maintained. 

Arnaud (1) thinks that the fertile stroma arises by the pro- 
liferation of a cell or a small number of cells which put out 
filaments extending radially on the cuticle of the host, and which 
fork enough to form a continuous bed. These proliferations 
constitute a regular radiate disk. As this plate becomes thick- 
ened on its inferior face, the superior face conserves on all its 
extent the radiate structure. 

Von Héhnel (16) states that the shield-shaped structure is 
merely a protective cover for the independent perithecia which 
develop inversely beneath it. 

Raciborski (7) was the first, however, to state that the peri- 
thecia were inverse. Theissen, Miss Doidge, Arnaud, and von 
Hohnel have accepted his statement as being correct. By’ in- 
verse, one understands the perithecia to be fastened on the under 
side of the hypha. The oldest or basal part therefore is above 
the newest or marginal part of the fruiting body. Von Hodhnel 
also pointed out that the asci arise from the part of the peri- 
thecium pressed against the leaf, or from the youngest and the 
top part of the structure. The asci are, therefore, inverted in 
the perithecium, but are borne upright on the surface of the leaf. 
This meaning of the term inverse has been used in this discussion. 

As I have stated previously, in Melzola the fructifications are 
always derived from hyphopodia, while in the Microthyriaceae 
in certain instances the perithecia are formed from the hypho- 
podium; in other instances from the cells of the mycelium. 
This latter method is characteristic of species which have a non- 
hyphopodiate mycelium, but which form as many fructifications 
as the others. 

The origin directly from the mycelium predominates through- 
out all the genera of the family. Out of the ninety-four species 
studied eighty-four show this method of development. 

Portions of the mycelium were fixed in cellodin, mounted and 
studied under oil immersion. The mycelium sometimes was 
slightly swollen in regions where a perithecium was developing. 


104 MYCOLOGIA 


At other times it remained normal. In either case, however, 
cross septa appeared, giving rise to one or more small cells. 
Rarely does the number exceed five. These cells are nearly 
square in shape, otherwise showing no differentiation from the 
rest of the mycelium. They all seem to function alike in the 
formation of the perithecium (PLATE 12, Fic. 8). 

They give rise to bud-like growths (PLATE 12, Fic. 4). These 
budded cells are irregular in outline, often distinctly lobed on 
their free margins. They may occur on both lateral margins 
of the mycelium. ‘The budded cells likewise divide increasing 
the number of cells centrifugally. These cells are united to one 
another so that a disk-like structure is produced (PLATE 12, Fic. 
6). Further cells are added to the structure until finally the 
maximum growth of the perithecium is reached. ‘The walls of 
these cells are thick and dark. There may be more than one 
layer of them. Cross sections of the perithecium generally show 
at least two. 

The second method of origin is from hyphopodia which are 
one- to two-celled structures located on the lateral margins of 
the mycelium. If the hyphopodium has only one cell, this cell 
buds several times (PLATE 14, Fic. 43). The budded cells in 
turn proliferate, the new cells being laid down in rows (PLATE 
14, Fic. 43). This process continues until the maximum growth 
has been reached. The radiate structure can be observed after 
several rows of cells have been formed. 

If the hyphopodium is a two-celled structure, the terminal 
cell only participates in the formation of the perithecium. This 
cell buds in the same manner as does the one-celled hyphopodium. 
In both kinds of hyphopodial development, as well as in the 
development from the mycelium, the perithecium produced is 
always inverse at maturity. 

A third type of development is from short lateral branches. 
This method is less common; only six species of all those investi- 
gated show this method of origin. The lateral branch of the 
mycelium, consisting of one to three cells, is ordinarily borne at 
an acute angle (PLATE 12, Fic. 14). The terminal cell in all 
cases is the one that functions in the production of the fruiting 
structure. This cell buds in a manner like that of the terminal 


RYAN: PERITHECIA IN MICROTHYRIACEAE 105 


cell of the hyphopodium. Asterina Gouldiae is a typical example 
of this mode. 

Only two species having nodulate cells in the mycelium were 
studied. These were Asterina inaequalis var. nodulosa (PLATE 
12, Fic. 13) and Asterina Schroetert (PLATE 13, Fic. 15). In both 
these forms the perithecium did not develop from the nodulate 
cell, but from an undifferentiated cell of the mycelium. The 
manner of growth was like that already described in the first 
method. 

Gaillard (13) gives illustrations of Asterina stricta which has 
nodulate cells that develop into the perithecia. This would 
indicate that there were two ways of producing a perithecium in 
the species which possess this kind of a mycelium. A further 
study of other species with a like mycelium would be interesting 
and would enable one to form a more accurate idea of the true 
method of origin in these species. 

In all the genera after the disk has reached its specific extension 
the radial growth is halted. The central part has meanwhile 
swollen from the middle of the perithecial hollow, while the 
peripheral growth remains close to the leaf. The result is a con- 
cave perithecium, which is only half a one in reality and is 
developed below the mycelium. 

The preceding description of the development of the peri- 
thecium can be applied to all the genera, for even those having 
oblong perithecia at maturity are at first circular. The elongate 
character is due to the more rapid growth of the perithecium on 
two opposite points on the disk. Frequently a three-angled 
perithecium is observed. This seems to be the result of growth 
at three points on the disk instead of at two. 

Contrasted to these various methods of origin, the perithecia 
in Meliola exhibit an unexpected constancy. In the forty species 
studied, all showed origin from the hyphopodia. Furthermore, 
the method of development observed agreed -in detail with that 
reported by Marshall Ward (17). A brief summary is given here. 

The simple pyriform body, the hyphopodium, after becoming 
swollen, suffers division into two portions or cells by a septum, 
usually vertically to the plane of the mycelium and leaf, and 
passing diagonally across the cavity with a slight curve so as to 


106 MYCOLoGIA 


abut on the outer walls at right angles, ornearly so. The original 
unicellular protuberance becomes in this manner divided into 
two more or less unequal cells which have different destinies. 
The more apical cel] divides more slowly and forms a mass of 
cells, the central ascogenous tissue, of the young perithecium. 
The outer cell divides much more rapidly, producing a layer of 
cells which gradually develops the thin-walled cells of the outer 
portion of the perithecium. These thin-walled cells make the 
ascogonium. In later stages, according to Ward, certain of the 
constituents are seen to form asci and spores, while others 
deliquesce and serve as nutritive material. The outer walls, 
as in the Microthyriaceae, become thick, hard, and dark colored. 
The perithecium in Meliola is always produced from the upper 
side of the mycelium so that it is never inverse (PLATE 15, Fic. 1) 
and is always spherical or approximately so. 

While this method of development holds for the majority of 
the species of Meliola, there are six species which show more 
distinct relationship to the Microthyriaceae. Beeli in his mono- 
graph of the genus Melzola (3) makes a division on the character 
of the radiate and pseudoparenchymatous tissue, placing the 
six species referred to in one group. He seems to have either 
overlooked the previous work of Theissen on Melzola (14) or to 
have rejected it. 

Theissen studied specimens of Meltola asterinoides and created 
for it a new genus, Amazonia, which he placed in the Micro- 
thyriaceae because of its radiate inverse perithecia. In studying 
material labeled Meliola Psychotriae, one of the other five species 
in this group, I have found that the development of the fruiting 
body approximates the development of the perithecium in the 
Microthyriaceae. The origin is from the hyphopodium, which 
proliferates in the manner already described for the hyphopodial 
origin in the Microthyriaceae. There is no definitely arranged 
method of division, as in Melzola, for the cells formed may or 
may not be of the same size. Furthermore, there may be more 
than two cells produced. In any circumstance the cells do not 
seem to form definite regions of the perithecium as in Meliola 
(PLATE 12, Fic. 7). At maturity or during the latter part of 
growth, the fruiting structure is inverse and radiate, thus differing 


RYAN: PERITHECIA IN MICROTHYRIACEAE 107 


in two essential characters from Meliola. For these reasons I 
have followed Theissen’s classification and placed the fungus in 
the genus Amazonia. 

Meliola Lagunculariae, like the foregoing species, is also inverse 
radiate (PLATE 15, Fic. 15). The method of development of 
the perithecium is like that described above. Therefore, this 
species should also be placed in the genus Amazonia as Amazonia 
Lagunculariae comb. nov. 

I did not study the other species listed by Beeli in his first 
division, but in all probability they will show the same characters 
as the two foregoing species and, therefore, will be placed in the 
same genus. 

In some of the Microthyriaceae an ostiole-like opening appears. 
This is not an “‘ostiole”’ in the true sense of the word. It appears 
at maturity and is due to the gelatinization or cracking of the 
central cells around the point of insertion on the mycelium or 
hyphopodium. The region where this occurs is the oldest and 
basal portion of the perithecium, and not the youngest and top 
part as one would ordinarily expect. There is evidently no pre- 
cise method of development for the “‘ostiole.’? When the peri- 
thecium is round the opening may be circular with ragged edges, 
due to the loss of several of the central cells (PLATE 14, Fic. 40), 
or it may be stellar, due to the severance of some of the cells 
along the lateral walls (PLATE 14, Fic. 33). Frequently these 
fissures extend almost to the margin of the perithecium. Or the 
entire central region of the perithecium may become gelatinous 
and disappear, leaving the asci fully exposed. 

In genera where the perithecia are oblong in shape, the 
‘“‘ostiole’’ follows the shape of the fruiting structure. It may 
remain narrow, so that it is thread-like in appearance, or it may 
become greatly widened, in which case the asci are exposed. In 
these oblong perithecia the lenticular ‘“‘ostiole’’ may become 
stellar at its ends. 

‘‘Ostioles,’’ however, do not occur in all the species before 
they reach maturity. Before the dispersal of the spores can take 
place though, the shield-like structure must be ruptured or 
become disintegrated. In Meliola, the rupturing of the peri- 
thecium occurs in the youngest and the top part of the peri- 


108 MYCOLOGIA 


thecium. This is diametrically opposed to that of the Micro- 
thyriaceae. In Amazonia Psychotriae the ‘‘ostiole”’ is like that 
of the family to which it belongs. 

The typical young fruiting body in the Microthyriaceae is 
always light brown in color. The radiate cell structure is ap- 
parent. Onmaturity it often becomes a dark brown, darker than 
the mycelium. The margin, though, remains lighter in color 
and the radiate character persists. At times, though, the 
radiate structure is not apparent, until the material has been 
boiled in potassium hydroxide, or a young specimen has been 
observed. When the first cells of the perithecium are laid, they 
ordinarily are arranged in a row. The next cells also occur in 
a row, and it is not until then that the perithecium begins to 
look radiate. On maturity, too, this character may be nearly 
obscured. In some genera the central cells gelatinize, so that 
only the marginal cells remain intact. These, however, always 
show the character. Other genera have perithecia which become 
almost wholly carbonaceous. But in such cases the margin 
again gives the character. 


SUMMARY 


The Microthyriaceae show four characteristic ways of develop- 
ing the perithecia. The methods are: from a cell of the my- 
celium; from hyphopodia; from the short lateral branch; from 
a nodulate cell. The most common way is from a cell of the 
mycelium. 

The perithecia are flattened. 

In early stages the fruiting structure is radiate. This char- 
acter later may be lost due to the perithecia losing its top through 
becoming gelatinous or carbonaceous. However, at all times the 
edge of the structure will show the character. 

The perithecia are all inverse, that is, they are borne on the 
lower surface of the mycelium. 

At maturity the perithecia break open, thus permitting the 
spores to escape. Often, however, an “‘ostiolar’’ opening appears 
at the base of the perithecium. This may be round, stellar, or 
linear in shape. 

In Meliola the perithecia always arise from hyphopodia. 


RYAN: PERITHECIA IN MICROTHYRIACEAE 109 


The perithecia are more or less round, never flattened. They 
never show the radiate character at maturity, although a few 
species may do so when very young. 

The perithecia are never inverse, but are always borne on the 
upper surface of the hyphopodium. 

- Meliola Psychotriae is radiate and inverse as is also Meltola 
Lagunculariae; they, therefore, belong to the genus Amazonia. 

An undetermined perisporiaceous form also showed the radiate 
perithecia and so rightfully belongs to Microthyriaceae. 

The following table lists the genera of the Microthyriaceae 
examined in this study. Hypho. stands for hyphopodia; myc., 
mycelium; lat. br., lateral branch; nod., nodulate cell. 


Method of Development Ostiole 
Genus Rt 8 ea wl See So SY ee th So le a ee 
Myc. |Hypho.| Lat. Br. | Nod. Linear Stellar |Round 
PUSTOVIN TG fase. 35 3 2 2 7 th 11 
Aisterinellas 02s 6 1 1 4 7 
Amazonia........ 1 1 i! 
Aulographum..... 1 1 1 
Calothyrio peltis.... 3 1 
PLEMTGNOGES fs es as: 3 3 
Echidnodella...... 5 5 
Englerulaster...... je 1 3 
TCTROOSTO aii a ees 10 10 
Morenoella........ 16 16 
Oestieria y Saal nd - 1 1 3 
Method of Development Perithecia 
Not Not 
Hypho. Inv. nee Rad. Rod 
MACHOL NOT, sah arin 40 2 38 4 36 


LITERATURE, CLEBD 


1. Arnaud, G. Les Asterinées. Théses présentés a la Faculté des Sciences 
de Paris. Série A, no. 805, no. d’ordre 1598. 1918. 

2. Atkinson, G. F. Phylogeny and relationships in the Ascomycetes. Ann. 
Missouri Bot. Gard. 2: 315-376. 1915. 

3. Beeli, M. Note sur le Genre Meliola Fr. Bull. Jardin Brux. 7: 89-160. 
1920. 

4. Doidge, E. M. South African Microthyriaceae. Trans. Roy. Soc. So. 
Afr. 8: 235-282. 1920. 

5. Gaillaird, A. Note sur les hyphopodies mycéliennes et la formation des 


périthecés des Asterina. Bull. Soc. Myc. Fr. 9: 95-97. 1893. 
6. Maire, R. Les sucoirs des Meliola et des Asterina. Ann. Myc. 6: 124- 


128.1908: 


110 MYCOLOGIA 


7. Raciborski, M. Paras. Algen und Pilze Java’s. 3: 43. 1900. 

8. Ryan, R.W. The Microthyriaceae of Porto Rico. Mycologia 16: 177- 
196. 1924. 

9. Saccardo, P. A. Syll. Fung. 2: 658. 1883. 

10. Stevens, F. L., & Ryan, R. W. Microthyriaceae [of Hawaii]. Bull. 
Bishop Mus. Honolulu 19: 66-78. 1925. 

11. Thaxter, R. Contributions toward a monograph of the Laboulbeniaceae. 
Mem. Amer. Acad. II. 12: 187-429. 1896; 13: 217-469. 1908. 

12. Theissen, F..& Sydow, H. Synoptische Tafeln. Ann. Myc. 15: 389-491. 
1917. 

13. Theissen, F. Ueber Membranstructuren bei den Microthyriaceen als 
Grundlage fiir den Ausbau der Hemisphaeriales. Myc. Centralbl. 3: 
273-286. 1913. 

14. ——. Ueber einige Mikrothyriaceen. Ann. Myc. 11: 493-511. 1913. 

15. ——. Hemisphaeriales. Ann. Myc. 11: 468-469. 1913. 

16. Von Hohnel. Ueber die Perithecien der Microthyriaceen und die Gattung 
Meliola Fries. Ber. Deuts. Bot. Ges. 35: 698-703. 1917. 

17. Ward, H.M. Onthe morphology and the development of the perithecium 
of Meliola, a genus of tropical epiphyllous fungi. Phil. Trans. Roy. 
Soc. London 34: 583-599. 1883. 


EXPLANATION OF PLATES 


All figures were drawn from material mounted in celloidin and drawn with 
the aid of a Spencer camera lucida, and were reduced one half in reproduction. 
PLATES 12 to 15 show stages in the development of the perithecia in the various 
species studied. 

PVATEE LZ 

Fig. 1, Morenoella decalvans var. Stigmatophylli; 2, M. Calami; 3, Asterinella 
multilobata; 4, A. Hippeasiri; 5, A. Ixorae; 6, A. Phoradendri; 7, Amazonia 
Psychotriae; 8, Calothyriopeltis Scaevolae; 9, Lembosia Sclerolobii; 10, L. 
Coccolobae; 11, L. ananensis; 12, L. portoricensis; 13, Asterina inaequalis var- 
nodulosa; 14, A. Gouldiae. 

PLATE 13 

Fig. 15, Asterina Schroetert; 16, A. acanthopoda; 17, Morenoella decalvans 
var. Rondeletiae; 18, M. Pothoide: var. Laevigatae; 19, M. Psychotriae; 20, 
M. decalvans var. Laugeriae; 21, M. Laugeriae; 22, M. miconicola; 23, M. 
impetiolaris; 24, M. Miconiae; 25, M. Melastomacearum; 26, Asterina dtplo- 
carpa; 27, A. Arnaudia; 28, A. Chrysophylli; 29, A. dilabens; 30, A. Rickit; 
31, A. Clermontiae; 32, A. Lobeliae; 35, A. dubit. 


PLATE 14 


Fig. 33, Asterina Ildefonsiae; 34, A. Fawcetti; 36, A. Hippocrateae; 37, A. 
correacola; 38, A. chrysophylliella; 39, A. kauaiensis; 40, Englerulaster 
orbicularis; 41, E. Papawiae; 42, Aulographum culmigenum; 43, Unidentified 


Perisporiaceae. 
PUATE a) 


Fig. 1, Meliola irregularis; 2, M. Lyoni; 3, M. Kaduae; 4, M. Koae; 5, 
M. maricaensis; 6, M. bidentata; 7, M. longipoda; 8, M. brasiliensis; 9, 
M. Dieffenbachiae; 10, M. Calophylli; 11, M. Juddiana; 12, M. hyptidicola; 
13, M. Cyperi; 14, M. Kaduae; 15, M. Lagunculariae; 16, M. cyclopoda; 
17, M. Marantae; 18, M. toruloidea; 19, M. brasiliensis. 


MYCOLOGIA VoLUME 18, PLATE 12 


DEVELOPMENT OF PERITHECIA 


VoLuME 18, PLATE 13 


MyYCoLoGIA 


oe 


you 


DEVELOPMENT OF PERITHECIA 


ne e aA ‘, 
- ¥ : At? 


a] 
: - 
* 
; 
e ‘ 
ys 
at 
, 
' 
‘ 
* 
a 
- 
= 
' 
‘ 
, 
) 
} 
j 
i 
. 

YT, 

Pendle | 

a 

} ‘ 
U i 

y « - 
. Cy 
. 

: 
« 
“ 


far wD Agee . 


* rogwet 4 J Ff eld 4 


MYCOLOGIA VoLUME 18, PLATE 14 


DEVELOPMENT OF PERITHECIA 


VOLUME 18, PLATE 15 


MYCOLOGIA 


DEVELOPMENT OF PERITHECIA 


Y 


AVS Rr 


Ruth Winifred Ryan was born at Appleton, Wisconsin, on 
June 19, 1899. She received her grade and high school education 
in the schools of that city. She entered Saint Clara College at 
Sinsinawa, Wisconsin, in 1917, and completed her undergraduate 
work with a major in botany in June, 1921, receiving the A. B. 
degree. She entered the University of Illinois in September, 1921, 
and began her graduate work in botany. She received the A. M. 
degree from this institution in June, 1923. She was graduate 
student in botany at the University of Wisconsin during the year 
1924-25. 


She was half-time Assistant in Botany at the University of 
Illinois for three years, where she taught general botany. 


She is a member of Sigma Xi, Sigma Delta Epsilon, and 
‘Theta Phi Alpha. 


Her publications are: The Morphology and Taxonomy of 
the Microthyriaceae of Porto Rico. Mycologia, 16:177-196. 
1924. “The Morphology and Taxonomy of the Microthyriaceae 
of Hawaii, by Stevens, F. L. and Ryan, R. W. Bulletin 10, 
Bishop Museum, 1925. A Systematic Presentation of New Genera 
Ota Dy ©) enteblnnkett. ba A. cL ouno) Ruth Wa kevane 
itansmamer.Micro. soce 32:43-67) - 19.23, 


3 0112 072902718 


AO i 


